Botanica Marina Vol. 45, 2002, pp.


© 2002 by Walter de Gruyter · Berlin · New York

A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia,

Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.

The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based

on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda

1998. – Taiwan: Anon. leg.Agardh to co-exist in the flora. – French Polynesia: Moorea. Sections were made using a freezing microtome. Nouméa. . Fig. P. New Caledonia. 3899). showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. a = axial cell. Tahiti. leg. Melanamansia glomerata. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. 11 Jul. 1998. 3. 1998. 3898. N’Yeurt (UPF 584). leg. leg. 1998. – Samoa: Palolo Deep Reserve. Prepared slides were examined using a Zeiss compound microscope.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). D. P. leg. a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. 1999. Kagoshima: 9 August 1959. M. Fiji. A. 2 Jul. The results of these investigations are described below. it was decided to revise records in the light of recent findings. Baba (SAP 056163). 25 October 1984 (SAP 046978). Photomicrographs were obtained using an Olympus BX-50 compound microscope. Sakai and S. A. 2 Jul. A. Tahiti. leg. Rintels (SUVA-A 3891. developed and printed in the laboratory. Skelton and Dave [SUVA-A 1565 (holotype). E. U. com. Tanegashima Island. Amansia rhodantha (Beqa. Okinoerabujima Island (60 km north of Okinawa). 1997. Okidomari. deep algal rim. leg. Punaauia. and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. Fig. 19 Mar. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. 2. Most recently. C. SUVA-A 5438L). – Cook Islands: Ngatangiia. Baba (SAP 056160). SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). Skelton. leg. 9 Feb. P. Rarotonga. N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. 18 Sep. Fig. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. Fiji. leg. Enomoto (SAP). 1. 3892. Figs 1–3. Payri (8/84. 5481L (spermatangial)]. Amansia rhodantha and Amansia glomerata. Material and Methods Specimens were collected by reef-walking. Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. A. A. leg. A. Furusato. Skelton and Dave (SUVA-A 1567). A. Y. N’Yeurt (SUVA-A 5128). 6 Jun. SUVA-A 5436L). R. cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell. 1989. 1–5 = order of formation of pericentral cells. snorkelling or SCUBA diving. ps = pseudo-pericentral cells). Afiti and D. pers..232 A. otherwise as in Holmgren et al. (1981). Tombleson and T. Skelton. M. 5480L (cystocarpic). 3896. in EPHE). 13 May 1982. Palolo Deep. and a camera-lucida attachment was used to make anatomical drawings.‘Al’ and ‘CG’ refer to specimens housed at IRD. SUVA-A 1566 (isotype). 3897). Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. 3895. Amansia rhodantha. 1984. leg. P. Taharaa. M. Faletose (SUVA-A 3894. 1998. N’Yeurt (UPF 347). 3893.

leg. Candelaria Zambales. leg. leg. Sumbawa. 3 and 5 March 1992.Revision of some Amansia and Melanamansia species 233 Table I. Basay. Kauai. Garrigue (SUVAA 5436L). 1983. leg.d. leg.Pagodpud.Palawan. C. Oahu. A. Eagles Nest. B79). Cordero and Tuangpalan (US 091395). G. Snellius-II (L 0108601). Agardh 1841: 26. H. Littler (US 163490). – Fiji: ‘Feejee’. Jul.. 3 and 15 Oct. leg. Oahu. 84 (L 0108603). Villeneuve (SUVA-A 1737). M. leg.d. leg. Yasawa Islands. SIPHILEXP-78 (US 013760). 1996: 472. leg.5 3. Ningaloo. (MURU. A. Figs 4–10 . N’Yeurt (SUVA-A 5437L). in TCD (Telfair. Ti Tree. Viti Levu. Kasahara (SUVA-A 1012. n. Kasahara (SUVA-A 1013. S.. leg. M. 1979. 13 July 1972. Chesterfield. 1984. 1985. leg. Garrigue (CH1). Isaac (L 0108595). 16 Nov. 10 Aug. 21 Sep.9 18. leg. n. Cannon (SAP 052062). M de Robillard (L 0108599). leg. 24 April 1996. Taxonomic results Amansia rhodantha (Harvey) J. J. Average values of selected characters for Amansia paloloensis. 1985.Canipo Is. Nanuyalevu. leg. Unia. Amansia rhodantha and Melanamansia glomerata surveyed in this study.2 53. R. – Fiji: ‘Feejee’. leg. C. 9 Sept. E. leg. – Kenya: Mombassa. Banilad. see text). A. leg. SIPHILEXP-78 (US 088530). Mauritius. 1984. LD 42600 / 0772). A. H. South (SUVA-A 5544. Rukuruku Village. R.Negros Oriental.W. 1980. n. Ovalau. 6 Jun 1978. 18 March 1992. Inhaca Peninsula. Jurien Bay WA. leg. M. 1994 (SUVA-A 797). S. Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39.P. C. 1838–42. – Philippines: Actin. J. P. – Australia: Heron Island QLD. Lopta. R127).Moreland (US 50851). 1965. Table I presents the average results of morphological measurements carried out on Amansia rhodantha. Wilkes Exploring Expedition (US 04057a). B. – Indonesia: Cape Batu Kerapo. 3 Feb. Naukathura Island. Lawai Kai. P.9 11. 25 Feb. Namada. Barrow Island. Plongeurs IRD (AL 288a. C. 2000: 188. Salayer. Masuda et al. Vatualailai. c. 1995. 1982. Bahati. Dumaguete.1 22. 29 May 1978. 1069). Ballou (SUVA-A 5438L). Hanauma Bay Beach Park. leg.leg. – Kenya: Mombassa. 1995. Snellius-II (L 0108602). SIPHILEXP-79 (US 088533). J. 3 Jun 1978. Agardh). 1954. leg. 1876. P. 17 Jul. SIPHILEXP-78 (US 088532).Dumaguete City. leg.Siquijor Is. leg. – Philippines:Gao-oa. leg. 1838–42. J. 27 Dec. Negros Oriental. Suva Reef. leg.. A. 10 May 1978.7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa. 1988. Potipot Is. G. R. Bay of Sanggar. leg. leg. Viti Levu. figs 1–11 (as Amansia glomerata C.Calindagan Reef. R. 1982.14 May 1979. 1067). Keats.Solong-On. Suva Barrier Reef. leg. E. B. AL 288b). Huisman (CWC 1997.. 84. leg. Huisman (MURU. figs 37–42. W. – Mozambique: Ponta Abril. leg. Magruder (US 086459). 1995. Norris 1988b: 211. 1981. Gaudichaud (‘Amansia glomerata’. South (SUVA-A 5431). JH240). Daida (US 086755). leg. G. R.23 May 1978. 28 Nov.. 30 Jun. Hildebrandt (L 0108600). leg. Carlson (SUVA-A 281). Richer de Forges (AL 288d-CG 77a).. Wilkes Exploring Expedition (US 04057b) (doubtful record. Beqa Lagoon. leg. M. Jul. NR 248). 1997. J.d. 1980. 15 Oct. Farlow (L 0108597). 1834). – New Caledonia: Lajeu Sudouest. leg. W. leg. 14 Feb. 26 Nov. 06 Sept. 347). B.Cuyo. Lectotype. Magnesia Virae. 14 March 1998. – New Caledonia: Lajeu Sudouest. B262). W. South in lit.. 21/22-Sept. L 0108598). leg. paloloensis and Melanamansia glomerata specimens examined in this study. (US 02890). – Lord Howe Island: 18 Aug. 25 September 1999. leg. 24 Nov. leg.). Cribb (SAP 051714). 16 Oct. Oahu. 17 Mar. Montebello Islands. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. Huisman (MURU. Kauai. W. Amansia rhodantha: n. Hildebrandt (L 0108596. fide G.d. 1973.Negros Oriental.Siaton.Ilocos Norte.7 54. CG 77b). leg. G. 84. 1819. Tangangge. SIPHILEXP-78 (US 088531). 5 June 1967. 1979. Yasawa Group. Catanduanes. Barrow Island. E. Villeneuve (SUVA-A 1774). N’Yeurt (SUVA-A 346. Hakula. leg. leg.3 2. leg.Malo. Isaac (L 0108594). A. D. Kipukai. anon.d. Viti Levu. leg. leg. SIPHILEXP-78 (US 088534). Huisman (MURU. Garrigue (CG 77c. leg. 10 July 1992. Skelton and G.. A. Huisman. South (SUVA-A 2460).11 Feb.6 3. Diani Beach. H. Hapmak. Cordero and Tuangpalan (US 091394). Silva et al. Negros Oriental. 16 March 1991. n. 1876.leg. J. 07 Oct. A. leg. leg. J. Tantabiddi.

Rehydrated type material of Delesseria rhodantha Harvey. US 02890 (scale bars = 3 mm). 8. Indian Ocean. from Rotuma Island. Surface view of segment cells of blade alae. Fig. N’Yeurt Figs 4–10. Fig. Habit of mature plant from the type locality. Fig. Figs 4. R. Amansia rhodantha. 9. SUVA-A 5437L. D.5 mm). 10. showing both lax and dense growth forms. South Pacific (scale bar = 10 µm). Type material of Delesseria rhodantha Harvey. showing axial cell (a) surrounded by five pericentral cells. Secondary branchlet. Fig. .234 A. 6. Type material of Delesseria rhodantha Harvey. Detail of tetrasporangial stichidia on lateral endogenous branch. 5. showing marginal serrations. in TCD (scale bar = 2. Axial structure of blade. in TCD (scale bar = 25 µm). 7. Mauritius. and a pair of budding branchlets from the midrib in the lower half. in TCD (scale bar = 200 µm).

5 mm). SUVA-A 5479L (scale bar = 1 mm). showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm). Fiji. Fig. 12. SUVA-A 1565 (scale bar = 25 µm). Note marginal cystocarps (arrowheads). Fig. showing lateral endogenous branches and midrib. Detail of the holotype (SUVA-A 1565). Holotype of Amansia paloloensis. 11.Revision of some Amansia and Melanamansia species 235 Figs 11–16. SUVA-A 3892 (scale bar = 12 mm). Detail of blades. 15. Compare with Fig. SUVA-A 3892 (scale bar = 2. 13. SUVA-A 5436L (scale bar = 50 µm). Tetrasporangial stichidia of Amansia rhodantha from Beqa. Fig. Habit of mature plant. with numerous rosette axes. Amansia paloloensis from Samoa. Fig. . 14. Segment cells of blade alae in surface view. Fig. 16. Fig. 4. Detail of morphologically distinct mature rosette and corticated axis.

Okamura 1900: 71. Fig. A central midrib is present. Blades of plant assumed to be from the Fiji Islands (see text). Isaac and Isaac 1968: 24. Apia. Individual tetrasporangia are spherical and tetrahedrally divided. Figs 11–16 Misapplied names: Amansia glomerata C. E. and are crisp and lanceolate.4. holotype in TCD. Plant from Potipot Island. 100–112 µm in diameter. CG77a (scale bar = 1 mm). Carposporangia are lanceolate. 0. Agardh 1824: 194. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). 25–42 × 83–125 µm. 23. Figs 17–24 Basionym: Amansia glomerata C. Yamada and Tanaka 1938: 86. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets.8–14. Tetrasporangial stichidia are pinnately arranged on either side of blade margins. L 0108598 (scale bar = 5 mm). showing smooth margins and elongate shape. Fig. pl. The main axis is irregularly branched. figs 2–14 (type locality: Palolo Deep Marine Reserve. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. in up to 10 regularly disposed pairs. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). N’Yeurt et al. 1995: 150. Axial structure of plant from Lajeu. Isaac 1956: 188. figs 19–21. 2. 77–110 µm thick. Plant from Mombassa. Japan. 1. East Africa. Kenya. 4B. with a range of 1–5. Note that the stichidia is twice laterally branched. Amansia paloloensis South et Skelton 1999: 247. and the apex is characteristically in-rolled. Womersley and Bailey 1970: 336.5 µm in diameter that lies adjacent to the axial cell (Figs 3. 21. Weber-van Bosse 1923: 369. AL288b (scale bar = 5 mm). Millar 1999: 523 (unverified record). plate CXXVI. Norris 1995: 67. Detail of blade of plant from Mombassa.236 A. fig. New Caledonia. and are initially vesiculate. Blade margins are smooth. to 0. or with regular serrations (endogenous branches). Philippines. up to 750 µm long and 250 µm broad. 17. – Melanamansia glomerata (C. e–i. Agardh 1824: 247. Secondary blades develop as successive pairs on the dorsal side of the midrib. N’Yeurt 1996: 428. Silva et al. XXV.8 mm in diameter. becoming narrower and disappearing towards the apex. 1. and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide. Internal structure is cellular. pl. 85–110 µm thick. containing up to 10 Figs 17–24. Norris 1995: 66) is usually 4. Kenya. New Caledonia. Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. New Caledonia. figs 14–29. Lewis and Norris 1987: 22. Garrigue and Tsuda 1988: 63. Plants are brownish-red. Blade of plant from Lajeu. Tetrasporangial stichidia of plant from Oahu. they are up to 4 cells long and non-vesiculate. L 0108598 (scale bar = 700 µm). fig. Abbott 1999: 404. with in-rolled leaf apex. 6E (unverified record). 1996: 87. Fig. Vietnam. South-East Asia. figs 1–13. Fiji. Mauritius. composed of two overlapping layers of elongate hexagonal cells in transverse rows. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. Millar et al. 19. N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. 198. Philippines. Payri and N’Yeurt 1997: 897. surrounded by five pericentral cells. Trichoblasts are sparse. 118C–J. R. Fig. Fig. fig. Melanamansia glomerata (C. with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. The stem of plants is thick and cartilaginous. Coppejans and Millar 2000: 333 (unverified record). 1999: 573. 31 fig. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. . Falkenberg 1901: 416. Known distribution: Hawaiian Islands. 18. Ultimate branchlets are ecorticate. stem-like and rigid in basal portions. New Caledonia. figs 127. Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. Deciduous.5–0. 185. Distribution: Tropical Indian Ocean. E. 20. 197. 436B. 19). 128. with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. D. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. 24. 267. Yoshida et al. 212a. Melanamansia glomerata. Kenya. Fig. up to 100 (on average 40) mm high. pl. The thallus is rose-red. Payri and Meinesz 1985: 511. fig. on average 53 mm high. Plant from Unia. each cell is about 20–20 × 87–100 µm in surface view. figs 1–11. 6. 22. Fig. 20. Tetrasporangia occur in flattened. fig. 1987: 61. Cribb 1983: 106. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. 9). SUVA-A 5431 (scale bar = 50 µm). Norris 1988b: 211. Jaasund 1976: 131. Agardh) R. Dublin). figs 1–3 (type locality: Cap Malheureux. Samoa). Norris 1995: 66) is on average 4. Norris 1995: 67. Tropical Pacific Ocean. Fig. US 091395 (scale bar = 5 mm). Agardh) R. Marginal teeth are up to 1 mm long and 445 mm broad. Kylin 1956: 544. 7 mm broad. Tsuda and Wray 1977: 103. Fritsch 1945: 570. pl. pl. US 4057b (scale bar = 1 mm). CG77a (scale bar = 10 µm). fig. up to 35 mm long and 6 mm broad. with a range of 3–6. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. The central axial cell is 28–30 µm in diameter. denuded below.

Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter. procarps and cystocarps occur on endogenous marginal branches. and stained the herbarium mounting paper. . the colour of the Melanamansia plants was noticeably brownish-red. Spermatangial heads are curved and arranged in marginal clusters. In all cases.

and absence of pseudopericentral cells. which is welldocumented and illustrated in the protologue of the species. M. rhodantha. rhodantha from Western Australia (MURU B262. rhodantha as described and illustrated by Norris (1988b: 211). regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent. More importantly. The Wilkes collection consists of two plants. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata. From this study. paloloensis. from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. Masuda pers. Kenya. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. thicker. In the Fiji Islands. serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. length of the blades. rhodantha as a pan-tropical entity. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. paloloensis from the type locality. Characters such as the degree of rosette formation. and Ceramium. Smith (1979: 38–40) points out that the accounting and . 12). and these authors considered the lack of highly corticated axes. 14) and corticated rosette axes (Fig. The present author has re-examined the holotype. Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted. one of which (US 04057a) is Amansia rhodantha. while the other (US 04057b) is Melanamansia glomerata. Now that the Samoan species has been compared with genuine herbarium material of A. rhodantha from the type collection and localities in Mauritius. com. isotype and other collections of A. loriform blades reminiscent of some specimens of A. Fiji. paloloensis showed no major difference in the range of characteristics among them. The drawing of the holotype (South and Skelton 1999: 246. Vidalia. 23) but the plant has not been collected in Fiji waters since.13). the Cook Islands and French Polynesia (Table I) with A. Rotuma. D. and a cross section of the blade clearly shows the five pericentral cells around the axial ring. MURU 248). New Caledonia. 2) indeed shows a plant with loriform. and trichoblasts lacking protective vesicles when young (Norris 1988b). plate CXXVI. Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. indicating that the morphology of the species is influenced perhaps by environmental factors. a comparison of material of A.). Fig. 3) which distinguishes that plant from A. Mozambique and Western Australia. and a weakly developed midrib as characters distinguishing A. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U. paloloensis from A. while Melanamansia becomes dark brown. On the other hand. with for instance A. rhodantha (South and Skelton 1999: 249). it appears to be conspecific with the latter species and thus extends the status of A. N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. a re-examination of the lectotype of Amansia glomerata C. the Philippines. the absence of blades organized in morphologically distinct tight axes around the parent axis. R.238 A. Amansia tends to bleach to translucent white. more coriaceous branchlets with less apparent midribs (this study). when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. The reproductive and trichoblast anatomy of A. rhodantha from the Indian Ocean and other localities. On the basis of these observations. as well as Amansia species from Taiwan. The habit and description of the holotype by Harvey (1834: 151. since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. an additional field test to distinguish Amansia spp. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. paloloensis. apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. These plants also had blades with well-developed midribs in the lower half (Fig. between 1838 and 1842 (US 04057b. However. Eastern Australia. generally smaller. Kuetzingia. having tight rosettes as well as very lax.S. Japan. fig. is comparable to A. While there is no doubt as to the generic identity of the latter specimen. Exploring Expedition. and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11.

13). rhodantha. environmental factors seem to play a role in determining the degree of rosette formation. calm waters (such as blue holes. which also had a consistently thicker thallus. When pseudo-pericentral cells were present. when present).e. in the case of A. Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. the description of the species states that branches have the same shape as the main axis (i. Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. it was observed that rosettes of blades along the secondary axes. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. the presence or absence of bladelets in rosettes. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a. and a main axis with alar cells not very distinct from secondary branches. both on the same plant and within individuals in similar populations. 1995). Norris (1988b: 211) reported that rosette axes in . type of endogenous branching. The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. an examination of the holotype and liquid-preserved and pressed isotypes of A. Indeed. and is backed by other morphological and biochemical differences (Norris 1988b. Indian Ocean Amansia rhodantha also lacked such cells. adding to the confusion between the two genera. degree of marginal serrations and thallus flattening. glomerata occurs in the same habitat as Amansia rhodantha (e. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24). rhodantha. L 0108603). fig. and would seem to function as attachment haptera. Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. These ventral. and hence the precise locality and date of this Melanamansia collection from Fiji is unknown. 19). except the Hawaiian samples. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. perhaps an adaptation to the exposed habitats where the plants came from. irregularly dichotomous proliferations lack alar cells. and in some instances lacking altogether. 7). fig. while in Malaysia Masuda et al. the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e. It would seem that perhaps shallow. 1996. some specimens were virtually indistinguishable from specimens of A. 1) shows both the presence of tight and lax rosettes of secondary branches. com. Fiji and New Caledonia (Table I). In most instances. pseudo-pericentral cells were universally absent. Payri and N’Yeurt 1997) were reexamined. Also. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13).g. For instance.Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. The illustration by Harvey of the holotype of Delesseria rhodantha (1834.). both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247. so it cannot be confirmed at this stage if M. fig. 5. more corticated secondary and primary axes with tight rosettes. paloloensis. pl. which was the main reason for separating the species from A. in New Caledonia and Kenya). Wilson and Kraft 2000). Masuda. Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific. older plants tend to have thicker. In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. they were clearly apparent and about half the size of pericentral cells (Fig. However. M. in some cases. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. M. paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. All Amansia rhodantha specimens examined had fairly prominent. From an examination of the specimens at hand. However. the Philippines. and found to lack pseudo-pericentral cells. and some samples from Kenya. Generally. 41) similarly report the absence of pseudo-pericentral cells in A. Ecological studies would be needed to verify these hypotheses. Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. The author has unsuccessfully tried to obtain material from the latter locality. in the case of Amansia paloloensis). (2000: 188. 2). rhodantha from Mauritius and Western Australia (compare Figs 4. thus placing them in Amansia. glomerata occurs in the Solomon Islands. alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig.g. CXXVI. Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined. which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight. while younger plants have less cortication and more lax or absent rosettes of blades. pers.

. These differences reinforce the conspecificity of the Amansia rhodantha and A. blade dimensions. as well as the generic distinctness of Melanamansia glomerata. and occur in two overlapping and slightly offset layers radiating on either side of the midrib. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A. in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. com. as well as more corticated axes. R. Melanamansia glomerata from all localities studied have consistently smoother. An examination of these elegant cells in species of Amansia (Figs 10. Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. Cover cells usually totally cover the developing sporangia. However. Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal. the number ranged from 3 to 7. only being more or less surrounded by cortication. it was noted that in Melanamansia the range was more restricted. which was also recorded by Norris (1988b) for Hawaiian material.4 (Norris 1995).240 A. An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades. B. in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. and found them relatively constant. rhodantha and A. One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig. showing the currently known distribution. Fig. This also varied according to the degree of rosette formation and age of the plant. paloloensis specimens studied. with most specimens having segment lengths of about 100 µm. which are in fact determinate branches covered by the pericentral cell derivatives. a number of preliminary conclusions can be made: A. rhodantha specimens observed in this study. with the length of segments ranging from 80 to 120 µm. as the axial ring was still discernible in all sections made. From Table I. usually 3 or 4 with an average of 3. which was not seen in Amansia rhodantha (Figs 6. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle). but can also occur apically.3. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. In A. There appears to be no notable difference in the range of thallus size. smaller and narrower blades. cortication of axes and rosette formation between specimens of Amansia rhodantha studied. Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. paloloensis. rhodantha studied. 16) and Melanamansia show no particular difference. 15). In Melanamansia it averages 4. 25.). but the range is reportedly less than in Amansia rhodantha. 20). as well as between A. This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. D.

pls v–vii. Berlin. P. South pers. E. Lund.). J. Intertidal Seaweeds in Tanzania. Masuda. R. Soc. First edition. University of Tromsø. J. 1841. W. East Africa Nat. 1945. 458 figs. Marine botany of the Kenya coast. 3. Norris is thanked for his helpful insights and suggestions for this research.A. Dr R. com. Isaac. II. H. 2. 27: 7–28. Kylin. Ms Susanna Riebe (LD – Botanical Museum. xvi + 477 + [1] pp. Mar. and R. K. flora and vegetation. Lewis. In historiam algarum symbolae. Bot. Sapporo). New Caledonia and the Philippines (this study). V. 1981. Observations sur la physiologie des algues marines. Jaasund. 1968. Professor G. S. xiv + 939 pp. 173 + (2) pp. xxxviii + 312 pp. J. Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Leiden). 1901. M. J. 29: 1–38. and A. Keuken and E. Accepted 17 January 2002. Washington). pls cxxv. 1988. G. General account of the environment. 1824. Smiths. Seven species of Rhodophyceae. Professor Willem Prud’homme van Reine (L – Rijksherbarium. iii + 159 pp. K. C. Kenya (this study). Myxophyceae. ser. 71 pls. Robin South and Mr Posa A. (2000) confirm that the genus Amansia does exist in the Pacific Ocean. J. Foreword. pigments reddish-brown. 1809. Acknowledgements The author sincerely thanks the following persons and institutions for the generous loan of type material and specimens used in this study: Dr John Huisman (MURU – Murdoch University). H. and J. Bull. and it appears that Melanamansia glomerata is so far restricted to Hawaii (Norris 1995). Garrigue. Die Gattungen der Rhodophyceen.. cxxvi.). J. Rhodophyceae. Marine Algae of the Southern Great Barrier Reef. N. P.A. The morphology and classification of some Ceramiaceae and Rhodomelaceae. Figure 25 presents a world distribution of these two genera as it is currently known. A history and annotated account of the benthic marine algae of Taiwan. M. E. trichoblasts on every second or third axial cell. Soc. Hommersand. Millar for his generous assistance in obtaining copies of rare publications. Lunda (Lund). S. S. 1956. Cal. Essais sur une classification des algues et des polypiers calcifères de Lamouroux. 43: 315–346. Gleerups Förlag. Bohn. Masuda pers. Honolulu. Marine algae of Inhaca Island and of the Inhaca Peninsula.. F. 1834. It is clear that both A. 2 folded maps. . Fritsch. Agardh. Nouv. Kawaguchi and S. Phil. 1963. E. Monographie 26. 2000. 24 pls. initially vesiculate and composed of up to 8 cells: Amansia rhodantha. Mr Barrett Brooks and Dr John Sims (US – Smithsonian Institution. Cambridge University Press. Norris. Taxonomic notes on marine algae from Malaysia. Kato. Mar. Scheltema & Holkema. I. 1976. 2000. Rhodophyta. 1999. ments rose-red. Mus. E. I. Catalogue of marine benthic algae from New Caledonia. trichoblasts non-vesiculate. Tsuda. Robin South is also thanked for his helpful comments on an earlier draft of this manuscript. B. (Hooker) 1: 147–157. Revised Key to the Species of Amansia and Melanamansia in the South Pacific – Presence of paired dorsal pericentral cells. xv + 673 pp. 1842. xvi + 754 pp.W. Decaisne.Revision of some Amansia and Melanamansia species 241 Conclusions The results of this study and those of South and Skelton (1999) and of Masuda et al. Linnaea 15: 1–50.Vietnam and Malaysia (M. rhodantha and Melanamansia glomerata are more widely distributed than previously thought. Annal. with descriptions of some new and little known species. com. Micronesica 21: 53–70. Bot. Charles Telfair.. A. Phang. Professors Michio Masuda and Tadao Yoshida (SAP – Hokkaido University. 43: 181–190. Bot. J. Falkenberg. Sci. 336 figs. and F. The Structure and Reproduction of the Algae. C. T. African Bot. J. Phaeophyceae. Nat. Index Herbariorum. Pub. Fauna und Flora des Golfes von Neapel. E. – Absence of paired dorsal pericentral cells. Nat. Dublin). Harvey. Millar. 443–457. K. W. Australian Coral Reef Society: Brisbane. 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